Download as pdf or txt
Download as pdf or txt
You are on page 1of 17

19/1/23, 22:31 Fungal States of Minds | bioRxiv

HOME| SUBMIT| FAQ| BLOG


| ALERTS / RSS| ABOUT
| CHANNELS
Search 
Advanced Search
bioRxiv posts many COVID19-related papers. A reminder: they have not been formally peer-reviewed and should not
guide health-related behavior or be reported in the press as conclusive.
New Results  Follow this preprint

Fungal States of Minds

Andrew Adamatzky, Jordi Vallverdu, Antoni Gandia, Alessandro Chiolerio, Oscar Castro, Gordana Dodig-Crnkovic
doi: https://doi.org/10.1101/2022.04.03.486900
This article is a preprint and has not been certified by peer review [what does this mean?].
 0  0  0  0  0  0  49

Abstract Full Text Info/History Metrics


 Preview PDF

Abstract

Fungal organisms can perceive the outer world in a way similar to what animals sense. Does that mean
that they have full awareness of their environment and themselves? Is a fungus a conscious entity? In
laboratory experiments we found that fungi produce patterns of electrical activity, similar to neurons.
There are low and high frequency oscillations and convoys of spike trains. The neural-like electrical
activity is yet another manifestation of the fungal intelligence. In this paper we discuss fungal cognitive
capabilities and intelligence in evolutionary perspective, and question whether fungi are conscious and
what does fungal consciousness mean, considering their exhibiting of complex behaviours, a wide
spectrum of sensory abilities, learning, memory and decision making. We overview experimental
evidences of consciousness found in fungi. Our conclusions allow us to give a positive answer to the
important research questions of fungal cognition, intelligence and forms of consciousness.

1. The Basic Nature of Fungi

Cognition and intelligence in nature is a topic of debate from multiple points of view. While nowadays it is
acceptable to speak about the cognition and intelligence within the biological kingdom Animalia, and

https://www.biorxiv.org/content/10.1101/2022.04.03.486900v1.full 1/17
19/1/23, 22:31 Fungal States of Minds | bioRxiv

even up to certain degree in Plantæ[80, 23], it is still controversial to discuss those capacities in other
lifeforms such as Fungi, Protista and Monera, the latest corresponding to single-celled organisms
without true nucleus (particularly bacteria) [85, 57]. However, the current move in research towards
basal cognition and intelligence shows how already unicellular organisms possess basal levels of
cognition and intelligent behaviour [43, 44, 45, 51, 52, 16, 11]. Further perplexity arises when
considering consciousness in living organisms. Humans are conscious, and some allow consciousness
in animals provided by a nervous systems. But other living creatures are typically considered not having
consciousness at all. Our focus is on fungi, remarkable organisms with surprising cognitive capacities
and behaviours which can be characterised as intelligent, and this article will argue that they possess a
level of basal consciousness.

Fungi dominated the Earth 600 million years before the arrival of plants [84, 20]. Even today the largest
known living organism in the world is a contiguous colony of Armillaria ostoyae, found in the Oregon
Malheur National Forest, and colloquially known as the “Humongous fungus”. Its size is impressive: 910
hectares, possibly weighing as much as 35,000 tons and having an estimated age of 8,650 years [71].
Furthermore, current estimations sum up a total of 3.8 million existing species of fungi, out of which only
120.000 are currently identified [35], representing a promising biotechnological tool-set from which
human kind has slightly scratched the surface.

Fungi have represented for humans ever since both an ally and a foe, in the first case serving to
produce fermented food and beverages (just to cite the most important one, Saccharomyces cerevisiae,
fundamental for bread, beer and wine) and in the second case, able to attack the same raw stocks and
generate famine and devastation (Puccinia graminis responsible for stem, black or cereal rust) [29].
They have also shown particular features, including interaction with the nervous system of parasitised
superior organisms, to induce them performing actions which are instrumental to further fungi
propagation. This is the case of Ophiocordyceps sinensis, also known with its Tibetan name yartsa
gumbu, an enthomopathogenic fungus parasitising ghost moths larvae, that is able to induce them
standing vertically under the soil surface, to facilitate spores spreading in spring times [86]. Similarly,
Ophiocordyceps unilateralis a complex of species also known as “zombie ant fungus” – surrounds
muscle fibres inside the ant’s body, and fungal cells form a network used to collectively control the host
behaviour, keeping the brain operative and guiding the ants to the highest points of the forest canopy,
the perfect place to sporulate [55].

By studying Fungi kingdom we can better hope to understand the origin of life[60] and evolution of
cognition, intelligence and consciousness as they gradually emerge from basal forms and up. But,
beyond all the extremely important biochemical mechanisms that make them possible, a fundamental
aspect in their organisation emerges: consciousness.

https://www.biorxiv.org/content/10.1101/2022.04.03.486900v1.full 2/17
19/1/23, 22:31 Fungal States of Minds | bioRxiv

2. Neuron-like spiking of fungi

Spikes of electrical potential are an essential characteristic of neural activity [47, 14, 68]. Fungi exhibit
trains of action-potential like spikes, detectable by intra- and extra-cellular recordings [75, 63, 2]. In
experiments with recording of electrical potential of oyster fungi Pleurotus djamor (Fig. 1(a)) we
discovered a wide range of spiking activity (Fig. 1(b)). Two types were predominant high-frequency
(period 2.6 min) and low-freq (period 14 min) [2]. While studying other species of fungus, Ganoderma
resinaceum, we found that most common width of an electrical potential spike is 5-8 min [4]. In both
species of fungi we observed bursts of spiking in the trains of the spike similar to that observed in central
nervous system [25, 42]. Whilst the similarly could be just phenomenological this indicates a possibility
that mycelium networks transform information via interaction of spikes and trains of spikes in manner
homologous to neurons. First evidence has been obtained that indeed fungi respond to mechanical,
chemical and optical stimulation by changing pattern of its electrically activity and, in many cases,
modifying characteristics of their spike trains [7, 8]. There is also evidence of electrical current
participation in the interactions between mycelium and plant roots during formation of mycorrhiza [17]. In
[30] we compared complexity measures of the fungal spiking train and sample text in European
languages and found that the ’fungal language’ exceeds the European languages in morphological
complexity.

https://www.biorxiv.org/content/10.1101/2022.04.03.486900v1.full 3/17
19/1/23, 22:31 Fungal States of Minds | bioRxiv

Figure 1:

Recording electrical activity of fungi. (a) Setup with an array of differential electrodes pairs. (b) A variety of patterns of spike trains.

In [3] we recorded extracellular electrical activity of four species of fungi. We speculated that fungal
electrical activity is a manifestation of the information communicated between distant parts of the fungal
colonies and the information is encoded into trains of electrical potential spikes. We attempted to
uncover key linguistic phenomena of the proposed fungal language. We found that distributions of
lengths of spike trains, measured in a number of spikes, follow the distribution of word lengths in human
languages. The size of fungal lexicon can be up to 50 words, however the core lexicon of most
frequently used words does not exceed 15-20 words. Species Schizophyllum commune and
Omphalotus nidiformis have largest lexicon while species Cordyceps militaris and Flammulina velutipes
have less extensive one. Depending on the threshold of spikes grouping into words, average word
length varies from 3.3 (O. nidiformis) to 8.9 (C. militaris). A fungal word length averaged over four
species and two methods of spike grouping is 5.97 which is of the same range as an average word
length in some human languages, e.g. 4.8 in English and 6 in Russian.

General anaesthetics in mammals causes reduction of neural fluctuation intensity, shift of electrical
activity to a lower frequency spectrum, depression of firing rates, which are also reflected in a decrease
in the spectral entropy of the electroencephalogram as the patient transits from the conscious to the

https://www.biorxiv.org/content/10.1101/2022.04.03.486900v1.full 4/17
19/1/23, 22:31 Fungal States of Minds | bioRxiv

unconscious state [56, 36, 38, 76]. In words, a rich spiking activity is a manifestation of consciousness,
and reduced activity of unconsciousness.

In [5] we demonstrated that the electrical activity of the fungus Pleurotus ostreatus is a reliable indicator
of the fungi anaesthesia. When exposed to a chloroform vapour the mycelium reduces frequency and
amplitude of its spiking and, in most cases, cease to produce any electrical activity exceeding the noise
level (Fig. 2). When the chloroform vapour is eliminated from the mycelium enclosure the mycelium
electrical activity restores to a level similar to that before anaesthesia.

Figure 2:

Reduction of spiking activity of Pleurotus ostreatus under influence of chloroform.

To summarise, in experimental laboratory studies of electrical activity of fungi we demonstrated that


fungi produce neuron-like bursts of spikes which are affected by general anaesthetics. These
phenomena indicate that fungi can posses the same degree of consciousness as creatures with central
nervous system do.

3. Fungal cognition

Once accepting the unity of such a big fungal biological structure as a single living entity, we need to
face a second challenge, that is, the anthropocentric bias [79] which sees consciousness as an
exclusively human capacity. This latter is the main cause for the lack of interest in cognition and
intelligence in minimal living systems. At this very moment we can affirm that intelligence is a property
extended across all living taxa, and that we can even talk about minimal consciousness, starting at
microbial level [54, 43, 44, 45, 51, 52, 16, 11]. The functional requests that make possible the existence
of such huge fungal colony are beyond the simple or automated addition of neighbouring cells, but
require a level of cooperation and informational communication that make necessary to ask for a

https://www.biorxiv.org/content/10.1101/2022.04.03.486900v1.full 5/17
19/1/23, 22:31 Fungal States of Minds | bioRxiv

mechanism that makes possible all these processes, could it be a form of consciousness? [53]. From a
phylogenetic perspective fungi provide the mechanisms for the existence of plant synapses [11], a
fundamental aspect for enabling plants complex information processing.

Our departure point is naturalistic and follows a simple idea: the biological explanations which can be
identified using a functionalist approach supervene on chemical mechanisms; consequently, any
approach to the emergence of informational minds must rely on such embodied factors. On the other
hand, social interactions modify this process, forcing us to consider the emergence of mind as the
coupling between single individual units and collective behaviour.

Our approach to the study of fungal minds is not a panpsychist one (attributing sentience to matter), but
is based on an informational processing model in which we identify the fungal self-awareness
mechanisms which provide an empirical foundation for the study of fungal minds. Two questions are
orienting our study: are fungi sentient? and…if so, could we talk about fungal collective consciousness?

From mycorrhizal relations, we know that fungi interact with plants roots and allow the existence of
mycorrhizal networks, used by plants to share or transport carbon, phosphorus, nitrogen, water, defense
compounds, or allelochemicals. Thanks to this network, plants regulate better their survival, growth, and
defence strategies. Such symbiotic relationship provides fungi carbohydrates, which are used
metabolically to generate energy or to expand their hyphal networks, generating therefore the collective
mycelium. And such mycelium can be considered the superstructure from which the collective fungal
consciousness emerges. As magisterially described by Stamets [77] (page 4): “The mycelium is an
exposed sentient membrane, aware and responsive to changes in its environment. As hikers, deer, or
insects walk across these sensitive filamentous nets, they leave impressions, and mycelia sense and
respond to these movements. A complex and resourceful structure for sharing information, mycelium can
adapt and evolve through the ever-changing forces of nature These sensitive mycelial membranes act
as a collective fungal consciousness.”

From an evolutionary point of view, mycelia are a clear example of cooperation, but also are used as a
cheating mechanism in relation to host plants [22]. Cheating, besides, decreases when high genetic
relatedness exists, a key point for sustaining multicellular cooperation in fungi [15]. We’ve considered
possible cheating actions of fungi towards their mycorrhizal hosts, but how can they form collective living
forms without cheating among themselves? The answer is related to the concept of allorecognition. The
ability to distinguish self from non-self is beneficial not only for self-preservation purposes [69] but also
for protecting the body from external menaces, like somatic parasitism [28] [66]. We’ve seen how fungi
are able to distinguish between themselves and others, and how several mechanisms allow them to
work in colonies, to establish symbiotic or parasitic relationships with other living systems. Their

https://www.biorxiv.org/content/10.1101/2022.04.03.486900v1.full 6/17
19/1/23, 22:31 Fungal States of Minds | bioRxiv

biochemistry allows them to adapt their actions to the informational variations of the surrounding
conditions, and requires a cognitive system able to adaptively manage such actions.

4. Consciousness as self-cognition

When observing cognitive and intelligent behaviour (adequate decision making, learning, problem
solving) of fungi we may ask whether some kind of consciousness enables their goal-directed behaviour,
where consciousness is the ability to make sense of the present situation. One can search for the
consciousness and its markers starting with humans and investigate its evolutionary origins in other
living organisms. Comparing humans with simpler living organisms it might be useful to make the
distinction between primary and higher order consciousness. With minimal modifications we can adapt
the notion of “primary consciousness in humans to describe “primary consciousness” (that corresponds
to “basal cognition” in other organisms including fungi and even unicellular organisms.

Over the centuries, consciousness has been a puzzling phenomenon despite all the efforts of the
scientists who tried to unveil its mysteries. Plant cognition and intelligence has been a matter of study for
hundreds of years, and still there is an ongoing controversy on its definition and functional extent [27].
There is a strong resistance and reluctance to acknowledge intelligence and cognitive capacities
(including degree of consciousness) in other living beings.

Fungi can store information and recall it [32]. Fungal memories are procedural, what is associated with
anoetic consciousness. Fungi show self-nonself recognition patterns. Fungi can navigate and solve
mazes looking for a bait. Fungi perceive their environment guiding themselves to sources of light or
higher oxygen concentrations. Self-nonself recognition, synchronous perception of light, nutrients,
gravity, or gas, moisture or other chemical gradients, only involves response to internal or external
stimuli, thus these cannot be quoted to support the affirmation that fungi are conscious or self-aware.
Such behaviours can be also encoded in a computer, which will respond accordingly to the instructed
parameters.

Fungus has a role in the managing of such aneuronal consciousness, as has been observed on tree
colonies[34]. The fundamental part of our debate is to show that living systems without nervous systems
(which we usually regard as necessary for self-awareness or intelligence) are able to perform tasks that
we usually would ascribe to conscious systems, and that consciousness is ubiquitous[81].

5. Biosensing for Data Integrating: the Mechanistic Path to Fungi Consciousness

https://www.biorxiv.org/content/10.1101/2022.04.03.486900v1.full 7/17
19/1/23, 22:31 Fungal States of Minds | bioRxiv

The first aim of consciousness, from a pragmatic point of view is that of collecting and combining
information at some specific level of detail in order to take a decision for action. Up to now the main
focus on consciousness studies has been focused on high level cognitive processes that follow a top-
down hierarchical structure. Using this model, fungi should be automatically discarded as suitable living
systems which could show consciousness. Instead of it our approach to the notion of consciousness will
follow a bottom up approach, from basic data to its ulterior processing and the possible conscious
decisions. We will present this case from a context situation: Mycelium typically is just under the humus
(the soil cover, i.e. a mix of leaves, needles from pines, fallen branches etc). Thus when we walk in the
forest mycelium “knows” by mechanical stimulation and stretch-activated receptors [21] that we are
walking there. This mechanoception process is shared by several living systems, including plants [58].

Consider for example calcium signal transduction [49]: it has been proven that in fungi this signalling
pathway has an essential role in the survival of fungi [88], as well as mediate stress responses, or
promote virulence [61]. Mechanosensitive channels can also be important for mating, as we see in
Neurosopora Crassa [46]. The existence of such mechanosensitive receptors in fungi make possible to
extend some cognitive properties we have already clearly defined in plants or mammals to fungi.
Consider for example the purinergic signalling [1]. Furthermore, sensing capabilities so far described
include also nutrient sensing (glucose, nitrogen) and general chemophysical sensing (pH, temperature,
light, gravity, electric field) [21]. Are those mechanisms a sufficient basis for the grounding of the
following questions: What does mycelium feel about this? Can the mycelium trace our movement? Can
the mycelium predict that we are approaching fruiting bodies (mushrooms)?

5.1. Integration data in fungi systems

All cognitive systems display mechanisms for using captured information (an active process, not just a
passive one) and to decide output actions. Due to the multimodal nature of data, this process implies an
integration and meaning hierarchies. The properties of fungi in data integration are shown in Tab. 1.

Table 1:

Different cognitive tasks performed by fungi.


5.2. The Self: from the genetic view to the perceptual and processing ones

It is necessary to say that the search for published scientific papers on the topics about “fungal
cognition” offers us zero results. The only connections between fungi and cognition are related most of
times to the cognitive impact for humans who are in contact with fungi. But, of course, fungi do perform
cognitive tasks, being self perception one of the most important.The most basic notion of self
identification is related to the delimitation of the structural elements that belong or not to one system. In

https://www.biorxiv.org/content/10.1101/2022.04.03.486900v1.full 8/17
19/1/23, 22:31 Fungal States of Minds | bioRxiv

this sense, fungi have shown that their hyphae are able to discriminate self/non-self and that use this
skill to decide to fuse themselves with other genetically compatible hyphae, thanks to anastomosis [70].
The mycelial networks of mycorrhizal fungi can also recognise correctly the roots of their hosts from
those of other surrounding non-hosts. Even different mycorrhizas can coexist but never fuse together.
There is a second mechanism implied in self recognition that is omitted in most of current studies:
alarmones [83]. These are regulatory molecules used to communicate exclusively among single cells
(not as host). Following [83], pages 13-14, we notice that a static organism, like a filamentous fungi can
live very long and move by hyphal growth, although it is physically constrained and must withstand the
onslaught of all potential genetic parasites they will encounter in their long life. The skill of self-
identification and colonial identity is therefore fundamental for several purposes (such as mating control).
And some fungi even use retro-parasites for their own development.

6. Discussion

We presented experimental laboratory and philosophical studies of the fungal states of mind. We
considered several aspects of fungal cognition and provided arguments supporting existence of fungal
consciousness. We raised many more questions than we provided answers. The new field of fungal
consciousness is opening in front of us. Let us discuss directions of future studies. How could we make
sure (in wording) that the fungal behaviour is not mechanical (automatic) responses but that it holds
intention? Abstraction, creativity, judgement, are characteristics of human consciousness. Are fungi able
of performing these? It might be useful to make the distinction between “primary” and “higher order”
consciousness.

Do fungi have holistic states of mind? Do they combine/modify such states? How many fungal states of
mind could be described? Do fungi create specific relational contexts? Or are they, on the contrary, not
capable of having holistic states of mind, ajust following completely prefixed patterns? At which extent
can we include fungi affects into such cognitive processing? Fungal chemotaxis could be part of such
proto-emotional states.

A deeper consciousness state allows us to understand and accept sacrificing our “self“ for higher
purposes (martyrs, heroes). Nevertheless we must recognise that human individuals (and animals as
well) are genetically different from each other, with the only exception of twins, while the same could not
apply to fungi. This in part supports the observation that fungi sacrifice their bodies for the sake of their
propagation. Another thought is about mortality: though it is, particularly in the western civilisation,
uncommon for people to live everyday life with a constant thought of being mortal, and pursue choices
that quite often go in the opposite direction, as if the individuals are immortal, it is unquestionably difficult

https://www.biorxiv.org/content/10.1101/2022.04.03.486900v1.full 9/17
19/1/23, 22:31 Fungal States of Minds | bioRxiv

for humans to figure out how the world would appear for immortal beings, like certain fungi. An immortal,
or even extremely old consciousness, would be able to develop perhaps an intelligence out of reach for
us, pursuing objectives that might seem unreasonable, for our limited perception. Perception of space
and time, causality, are all aspects that we consider our unquestionable bottom line. But given the
peculiarity of fungi morphology and degree of connection, we may imagine how radically different
computational schemes are embedded into a fungal consciousness. For example, rather than 3-
dimensional visual perception, holographic perception might be possible, considering the quasi-flat
distribution of mycelia and its mechanoceptive reconstruction of moving objects (animals) at the upper
boundary layer. Non-causal consciousness might arise from this specific perception framework,
eventually hindering the time perception. All of these remain open questions for further investigations.

7. Acknowledgement

AA and NP were supported by the funding from the European Union’s Horizon 2020 research and
innovation programme FET OPEN “Challenging current thinking” under grant agreement No 858132.

References

[1].↵ Maria P Abbracchio, Geoffrey Burnstock, Alexei Verkhratsky, and Herbert Zimmermann. Purinergic signalling in the nervous
system: an overview. Trends in neurosciences, 32(1):19–29, 2009. CrossRef PubMed Web of Science Google Scholar
[2].↵ Andrew Adamatzky. On spiking behaviour of oyster fungi pleurotus djamor. Scientific reports, 8(1):1–7, 2018. CrossRef
Google Scholar

[3].↵ Andrew Adamatzky. Language of fungi derived from electrical spiking activity. arXiv preprint arXiv:2112.09907, 2021.
Google Scholar

[4].↵ Andrew Adamatzky and Antoni Gandia. On electrical spiking of ganoderma resinaceum. Biophysical Reviews and Letters, pages
1–9, 2021. Google Scholar
[5].↵ Andrew Adamatzky and Antoni Gandia. Fungi anaesthesia. Scientific Reports, 12(1):1–8, 2022. Google Scholar
[6]. Andrew Adamatzky and Antoni Gandia. Living mycelium composites discern weights via patterns of electrical activity. Journal of
Bioresources and Bioproducts, 7(1):26–32, 2022. Google Scholar
[7].↵ Andrew Adamatzky, Antoni Gandia, and Alessandro Chiolerio. Fungal sensing skin. Fungal biology and biotechnology, 8(1):1–6,
2021. Google Scholar
[8].↵ Andrew Adamatzky, Anna Nikolaidou, Antoni Gandia, Alessandro Chiolerio, and Mohammad Mahdi Dehshibi. Reactive fungal
wearable. Biosystems, 199:104304, 2021. Google Scholar

https://www.biorxiv.org/content/10.1101/2022.04.03.486900v1.full 10/17
19/1/23, 22:31 Fungal States of Minds | bioRxiv

[9]. Mariana Cruz Almeida and Alexandra C Brand.Thigmo responses: the fungal sense of touch. Microbiology spectrum, 5(2):5–2,
2017. Google Scholar
[10]. Diana R Andrade-Linares, Stavros D Veresoglou, and Matthias C Rillig.Temperature priming and memory in soil filamentous
fungi. Fungal Ecology, 21:10–15, 2016. Google Scholar
[11].↵ Frantisek Baluska and Stefano Mancuso. Microorganism and filamentous fungi drive evolution of plant synapses. Frontiers in
cellular and infection microbiology, 3:44, 2013. Google Scholar
[12]. Jorge Barriuso, Deborah A Hogan,Tajalli Keshavarz, Mara Jess, and Jes Jess Martnez. Role of quorum sensing and chemical
communication in fungal biotechnology and pathogenesis. FEMS Microbiology Reviews, 022:627–638, 2018. Google Scholar
[13]. G. L. Barron and R. G.Thorn. Destruction of nematodes by species of pleurotus. Canadian Journal of Botany, 65:774, 4 1987.
CrossRef Google Scholar

[14].↵ Morris H Baslow.The languages of neurons: an analysis of coding mechanisms by which neurons communicate, learn and
store information. Entropy, 11(4):782–797, 2009. Google Scholar
[15].↵ Eric Bastiaans, Alfons JM Debets, and Duur K Aanen. Experimental evolution reveals that high relatedness protects
multicellular cooperation from cheaters. Nature communications, 7(1):1–10, 2016. Google Scholar
[16].↵ Eshel Ben-Jacob. Bacterial wisdom, gdels theorem and creative genomic webs. Physica A, 248:5776, 1998. Google Scholar
[17].↵ RLL Berbara, BM Morris, HMAC Fonseca, B Reid, NAR Gow, and MJ Daft. Electrical currents associated with arbuscular
mycorrhizal interactions. New phytologist, 129(3):433–438, 1995. Google Scholar
[18]. Lynne Boddy and T. Hefin Jones. Mycelial responses in heterogeneous environments: parallels with macroorganisms. Fungi in
the Environment, pages 112–140, 1 2007. Google Scholar
[19]. Alexandra Brand and Neil AR Gow. Mechanisms of hypha orientation of fungi. Current opinion in microbiology, 12(4):350–357,
2009. CrossRef PubMed Google Scholar
[20].↵Mark C. Brundrett. Coevolution of roots and mycorrhizas of land plants, may 2002. Google Scholar
[21].↵ D. Mailnder-Snchez C. Braunsdorf and M. Schaller. Fungal sensing of host environment, journal =. Google Scholar
[22].↵ JA Callow. Advances in botanical research.Vol 22. Elsevier, 1999. Google Scholar
[23].↵ Paco Calvo, Monica Gagliano, Gustavo M. Souza, and Anthony Trewavas. Plants are intelligent, heres how. Annals of Botany,
125(1):1128, 2020. Google Scholar

[24]. MJ Carlile.The photobiology of fungi. Annual Review of Plant Physiology, 16(1):175–202, 1965. CrossRef Google Scholar

[25].↵ JH Cocatre-Zilgien and F Delcomyn. Identification of bursts in spike trains. Journal of neuroscience methods, 41(1):19–30,
1992. CrossRef PubMed Web of Science Google Scholar
[26]. LM Corrochano and P Galland. Photomorphogenesis and gravitropism in fungi. In Growth, differentiation and sexuality, pages
233–259. Springer, 2006. Google Scholar
https://www.biorxiv.org/content/10.1101/2022.04.03.486900v1.full 11/17
19/1/23, 22:31 Fungal States of Minds | bioRxiv

[27].↵ Fatima Cvrckova, Helena Lipavská, and Viktor Žárský. Plant intelligence why, why not or where? Plant Signaling and Behavior,
4(5):394–399, may 2009. Google Scholar

[28].↵ Tamas Czaran, Rolf F Hoekstra, and Duur K Aanen. Selection against somatic parasitism can maintain allorecognition in
fungi. Fungal Genetics and Biology, 73:128–137, 2014. CrossRef PubMed Google Scholar
[29].↵ Kate MJ de Mattos-Shipley, Kathryn L Ford, Fabrizio Alberti, AM Banks, Andrew M Bailey, and GD Foster.The good, the bad
and the tasty: the many roles of mushrooms. Studies in mycology, 85:125–157, 2016. CrossRef Google Scholar
[30].↵ Mohammad Mahdi Dehshibi and Andrew Adamatzky. Electrical activity of fungi: Spikes detection and complexity analysis.
Biosystems, 203:104373, 2021. Google Scholar
[31]. Marina Fomina, Karl Ritz, and Geoffrey M. Gadd. Negative fungal chemotropism to toxic metals. FEMS Microbiology Letters,
193:207–211, 12 2000. CrossRef PubMed Web of Science Google Scholar

[32].↵ Yu Fukasawa, Melanie Savoury, and Lynne Boddy. Ecological memory and relocation decisions in fungal mycelial networks:
responses to quantity and location of new resources. ISME Journal, 14(2):380–388, feb 2020. Google Scholar
[33]. Kristi L Hanson, Dan V Nicolau Jr., Luisa Filipponi, Lisen Wang, Abraham P Lee, and Dan V Nicolau. Fungi use efficient
algorithms for the exploration of microfluidic networks. small , 2(10):1212–1220, 2006. CrossRef PubMed Google Scholar
[34].↵ John Laker Harley and JS Waid. A method of studying active mycelia on living roots and other surfaces in the soil.
Transactions of the British Mycological Society, 38(2):104–118, 1955. Google Scholar
[35].↵ David L Hawksworth and Robert Lucking. Fungal Diversity Revisited: 2.2 to 3.8 Million Species. Microbiology spectrum, 5(4),
2017. Google Scholar
[36].↵ Harald Hentschke, Cornelius Schwarz, and Bernd Antkowiak. Neocortex is the major target of sedative concentrations of
volatile anaesthetics: strong depression of firing rates and increase of gabaa receptor-mediated inhibition. European Journal of
Neuroscience, 21(1):93–102, 2005. CrossRef PubMed Web of Science Google Scholar
[37]. J. Hiscox, J. O’Leary, and L. Boddy. Fungus wars: basidiomycete battles in wood decay. Studies in Mycology, 89:117–124, 3 2018.
CrossRef Google Scholar

[38].↵ Axel Hutt, Jéeréemie Lefebvre, Darren Hight, and Jamie Sleigh. Suppression of underlying neuronal fluctuations mediates
eeg slowing during general anaesthesia. Neuroimage, 179:414–428, 2018. CrossRef Google Scholar
[39]. Saima Khalid and Nancy P Keller. Chemical signals driving bacterial-fungal interactions. 2021. Google Scholar
[40]. Ursula Kes,Weeradej Khonsuntia, Shanta Subba, and Bastian Drnte.Volatiles in communication of agaricomycetes. Physiology
and Genetics, pages 149–212, 2018. Google Scholar
[41]. Jonathan Leake, David Johnson, Damian Donnelly, Gemma Muckle, Lynne Boddy, and David Read. Networks of power and
influence: the role of mycorrhizal mycelium in controlling plant communities and agro-ecosystem functioning. Canadian Journal of
Botany, 82:1016, 8 2004. CrossRef Web of Science Google Scholar
[42].↵ CR Legendy and M Salcman. Bursts and recurrences of bursts in the spike trains of spontaneously active striate cortex
neurons. Journal of neurophysiology, 53(4):926–939, 1985. CrossRef PubMed Web of Science Google Scholar
https://www.biorxiv.org/content/10.1101/2022.04.03.486900v1.full 12/17
19/1/23, 22:31 Fungal States of Minds | bioRxiv

[43].↵ Michael Levin.The computational boundary of a self: Developmental bioelectricity drives multicellularity and scale-free
cognition. Frontiers in Psychology, 10:2688, 2019. Google Scholar
[44].↵ Michael Levin. Life, death, and self: Fundamental questions of primitive cognition viewed through the lens of body plasticity
and synthetic organisms. Biochemical and Biophysical Research Communications, 2020. In press. Google Scholar
[45].↵ Michael Levin, Fred Keijzer, Pamela Lyon, and Detlev Arendt. Uncovering cognitive similarities and differences, conservation
and innovation. Phil.Trans. R. Soc. B, 376:20200458, 2021. Google Scholar
[46].↵ Roger R Lew, Zohaib Abbas, Marinela I Anderca, and Stephen J Free. Phenotype of a mechanosensitive channel mutant, mid-
1, in a filamentous fungus, neurospora crassa. Eukaryotic cell , 7(4):647–655, 2008. Abstract/FREE Full Text Google Scholar
[47].↵ Michael S Lewicki. A review of methods for spike sorting: the detection and classification of neural action potentials.
Network: Computation in Neural Systems, 9(4):R53, 1998. Google Scholar
[48]. Edward D. Lipson. Phototropism in fungi. Biophysics of Photoreceptors and Photomovements in Microorganisms, pages 311–325,
1991. Google Scholar
[49].↵ Shuyuan Liu,Yinglong Hou,Weiguo Liu, Chunyan Lu,Weixin Wang, and Shujuan Sun. Components of the calcium-calcineurin
signaling pathway in fungal cells and their potential as antifungal targets. Eukaryotic cell , pages EC–00271, 2015. Google Scholar
[50]. Hong Luo, Minghe Mo, Xiaowei Huang, Xuan Li, and Keqin Zhang. Coprinus comatus: a basidiomycete fungus forms novel
spiny structures and infects nematodes. Mycologia , 96:1218, 2004. CrossRef PubMed Web of Science Google Scholar
[51].↵ Pamela Lyon.The biogenic approach to cognition. Cognitive Processing, 7:1129, 2005. Google Scholar
[52].↵ Pamela Lyon.The cognitive cell: Bacterial behaviour reconsidered. Frontiers in Microbiology, 6:264, 2015. Google Scholar
[53].↵ Lynn Margulis.The conscious cell. Annals of the New York Academy of Sciences, 929(1):55–70, 2001. PubMed
Google Scholar

[54].↵ Lynn Margulis, Celeste A. Asikainen, and Krumbein Wolfgang E. Chimeras and Consciousness: Evolution of the Sensory Self . MIT
Cambridge, 2011. Google Scholar
[55].↵Missy L. Hazen Raquel G. Loreto Colleen A. Mangold Danny Z. Chen Maridel A. Fredericksen,Yizhe Zhang and David P.
Hughes. Three-dimensional visualization and a deep-learning model reveal complex fungal parasite networks in behaviorally manipulated
ants, journal =. Google Scholar
[56].↵ P Miu and E Puil. Isoflurane-induced impairment of synaptic transmission in hippocampal neurons. Experimental brain
research, 75(2):354–360, 1989. PubMed Web of Science Google Scholar
[57].↵ Nicholas P. Money. Hyphal and mycelial consciousness: the concept of the fungal mind. Fungal Biology, 125:257–259, 4
2021. Google Scholar
[58].↵ Gabriele B Monshausen and Elizabeth S Haswell. A force of nature: molecular mechanisms of mechanoperception in plants.
Journal of Experimental Botany, 64(15):4663–4680, 2013. CrossRef PubMed Web of Science Google Scholar

https://www.biorxiv.org/content/10.1101/2022.04.03.486900v1.full 13/17
19/1/23, 22:31 Fungal States of Minds | bioRxiv

[59]. David Moore. Perception and response to gravity in higher fungi a critical appraisal. New Phytologist, 117:3–23, 1991.
CrossRef PubMed Web of Science Google Scholar

[60].↵ David Moore. Fungal biology in the origin and emergence of life. Cambridge University Press, 2013. Google Scholar
[61].↵ Eric M Muller, Nancy A Mackin, Scott E Erdman, and Kyle W Cunningham. Fig1p facilitates ca2+ influx and cell fusion during
mating of saccharomyces cerevisiae. Journal of Biological Chemistry, 2003. Google Scholar
[62]. Ronald No and E.Toby Kiers. Mycorrhizal markets, firms, and co-ops. Trends in Ecology Evolution, 33:777–789, 10 2018.
Google Scholar

[63].↵ S Olsson and BS Hansson. Action potential-like activity found in fungal mycelia is sensitive to stimulation.
Naturwissenschaften, 82(1):30–31, 1995. CrossRef Google Scholar
[64]. Joseph KE Ortega and RUSTEM Igor Gamow. Phycomyces: habituation of the light growth response. Science,
168(3937):1374–1375, 1970. Abstract/FREE Full Text Google Scholar

[65]. Robert M Page. Phototropism in fungi. Photophysiology: Current Topics, page 65, 2013. Google Scholar
[66].↵ Mathieu Paoletti, Sven J Saupe, and Corinne Clave. Genesis of a fungal non-self recognition repertoire. PLoS one, 2(3),
2007. Google Scholar
[67].THS Perera, DW Gregory, D Marshall, and NAR Gow. Contact-sensing by hyphae of dermatophytic and saprophytic fungi.
Journal of medical and veterinary mycology, 35(4):289–293, 1997. PubMed Google Scholar
[68].↵ J Andrew Pruszynski and Joel Zylberberg.The language of the brain: real-world neural population codes. Current opinion in
neurobiology, 58:30–36, 2019. CrossRef PubMed Google Scholar
[69].↵ Christian Q Scheckhuber, Andrea Hamann, Diana Brust, and Heinz D Osiewacz. Cellular homeostasis in fungi: impact on
the aging process. In Aging Research in Yeast, pages 233–250. Springer, 2011. Google Scholar
[70].↵ Hainfried EA Schenk, Reinhold G Herrmann, Kwang W Jeon, Norbert E Muller, and Werner Schwemmler. Eukaryotism and
symbiosis: Intertaxonic combination versus symbiotic adaptation. Springer Science & Business Media , 2012. Google Scholar
[71].↵ Craig L Schmitt and Michael L Tatum.The malheur national forest: Location of the world’s largest living organism (the
Humongous Fungus). United States Department of Agriculture, Forest Service, Pacific Northwest, 2008. Google Scholar
[72].Yanfang Shang, Peng Feng, and Chengshu Wang. Fungi that infect insects: Altering host behavior and beyond. PLOS Pathogens,
11:e1005037, 8 2015. CrossRef PubMed Google Scholar

[73]. Suzanne W. Simard. Mycorrhizal networks facilitate tree communication, learning, and memory, 2018. Google Scholar
[74]. Suzanne W. Simard, David A. Perry, Melanie D. Jones, David D. Myrold, Daniel M. Durall, and Randy Molina. Net transfer of
carbon between ectomycorrhizal tree species in the field. Nature 1997 388:6642, 388:579–582, 1997. CrossRef
Web of Science Google Scholar

[75].↵ Clifford L Slayman,W Scott Long, and Dietrich Gradmann. “Action potentials” in Neurospora crassa, a mycelial fungus.
Biochimica et Biophysica Acta (BBA)-Biomembranes, 426(4):732–744, 1976. Google Scholar
https://www.biorxiv.org/content/10.1101/2022.04.03.486900v1.full 14/17
19/1/23, 22:31 Fungal States of Minds | bioRxiv

[76].↵ James W Sleigh, D Alistair Steyn-Ross, Moira L Steyn-Ross, C Grant, and G Ludbrook. Cortical entropy changes with
general anaesthesia: theory and experiment. Physiological measurement, 25(4):921, 2004. CrossRef PubMed Web of Science
Google Scholar

[77].↵ Paul Stamets. Mycelium running: how mushrooms can help save the world . Random House Digital, Inc., 2005. Google Scholar
[78]. Darren D.Thomson, Silvia Wehmeier, Fitzroy J. Byfield, Paul A. Janmey, David Caballero-Lima, Alison Crossley, and Alexandra C.
Brand. Contact-induced apical asymmetry drives the thigmotropic responses of candida albicans hyphae. Cellular microbiology,
17:342–354, 3 2015. CrossRef Google Scholar

[79].↵ Anthony Trewavas. Plant behaviour and intelligence. OUP Oxford, 2014. Google Scholar
[80].↵ Anthony Trewavas.The foundations of plant intelligence. Interface Focus, 7, 6 2017. Google Scholar
[81].↵ Anthony J Trewavas and František Baluska.The ubiquity of consciousness:The ubiquity of consciousness, cognition and
intelligence in life. EMBO reports, 12(12):1221–1225, 2011. FREE Full Text Google Scholar
[82].Veronica Veses, Andrea Richards, and Neil AR Gow.Vacuoles and fungal biology. Current opinion in microbiology, 11(6):503–510,
2008. CrossRef PubMed Google Scholar
[83].↵ Luis P Villarreal. Origin of group identity: viruses, addiction and cooperation. Springer Science & Business Media , 2008.
Google Scholar

[84].↵ Daniel Y.-C.Wang, Sudhir Kumar, and S. Blair Hedges. Divergence time estimates for the early history of animal phyla and
the origin of plants, animals and fungi. Proceedings of the Royal Society of London. Series B: Biological Sciences, 266(1415):163–171, jan
1999. CrossRef GeoRef PubMed Google Scholar
[85].↵ Hans V.Westerhoff, Aaron N. Brooks, Evangelos Simeonidis, Rodolfo Garca-Contreras, Fei He, Fred C. Boogerd,Victoria J.
Jackson,Valeri Goncharuk, and Alexey Kolodkin. Macromolecular networks and intelligence in microorganisms. Frontiers in
Microbiology, 5:379, 2014. Google Scholar
[86].↵ Yi-Jian Yao Xiao-Liang Wang. Host insect species of Ophiocordyceps sinensis: a review, journal =. Google Scholar
[87]. Gillian Young. Fungal communication gets volatile. Nature Reviews Microbiology, 7(1):6–6, 2009. Google Scholar
[88].↵ Qilin Yu, Fan Wang, Qiang Zhao, Jiatong Chen, Bing Zhang, Xiaohui Ding, Hui Wang, Baopeng Yang, Guangqing Lu, Biao
Zhang, et al. A novel role of the vacuolar calcium channel yvc1 in stress response, morphogenesis and pathogenicity of candida
albicans. International Journal of Medical Microbiology, 304(3-4):339–350, 2014. CrossRef PubMed Google Scholar
[89]. Zhenzhong Yu and Reinhard Fischer. Light sensing and responses in fungi. Nature Reviews Microbiology, 17(1):25–36, 2019.
CrossRef Google Scholar

 Back to top
 Previous Next 
Posted April 03, 2022.
 Download PDF  Email
 Print/Save Options  Share
https://www.biorxiv.org/content/10.1101/2022.04.03.486900v1.full 15/17
19/1/23, 22:31 Fungal States of Minds | bioRxiv

 Citation Tools
Tweet Like 84

COVID-19 SARS-CoV-2 preprints from


medRxiv and bioRxiv
Subject Area

Animal Behavior and Cognition

Subject Areas

All Articles

Animal Behavior and Cognition


Biochemistry
Bioengineering
Bioinformatics
Biophysics
Cancer Biology
Cell Biology
Clinical Trials*
Developmental Biology
Ecology
Epidemiology*
Evolutionary Biology
Genetics
Genomics
Immunology
Microbiology
Molecular Biology
Neuroscience
Paleontology
Pathology
Pharmacology and Toxicology
Physiology
Plant Biology
Scientific Communication and Education
Synthetic Biology
https://www.biorxiv.org/content/10.1101/2022.04.03.486900v1.full 16/17
19/1/23, 22:31 Fungal States of Minds | bioRxiv

Systems Biology
Zoology
* The Clinical Trials and Epidemiology subject categories are now closed to new
submissions following the completion of bioRxiv's clinical research pilot project and
launch of the dedicated health sciences server medRxiv (submit.medrxiv.org). New
papers that report results of Clinical Trials must now be submitted to medRxiv. Most new
Epidemiology papers also should be submitted to medRxiv, but if a paper contains no
health-related information, authors may choose to submit it to another bioRxiv subject
category (e.g., Genetics or Microbiology).

https://www.biorxiv.org/content/10.1101/2022.04.03.486900v1.full 17/17

You might also like